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RESEARCH ARTICLE
Variation in Foot Strike Patterns among
Habitually Barefoot and Shod Runners inKenya
Daniel E. Lieberman1*, Eric R. Castillo1, Erik Otarola-Castillo1, Meshack K. Sang2, Timothy
K. Sigei3, Robert Ojiambo2, Paul Okutoyi4, Yannis Pitsiladis5
1 Department of Human Evolutionary Biology, Harvard University, Cambridge, Massachusetts, United
Statesof America, 2 Medical Physiology Department, School of Medicine, Moi University, Eldoret, Kenya,
3 Department of Statistics and Computer Science Moi University, Eldoret, Kenya, 4 Department ofOrthopaedics and Rehabilitation, School of Medicine, Moi University, Eldoret, Kenya, 5 Centre for Sport and
Exercise Science and Medicine, University of Brighton, Brighton, United Kingdom
Abstract
Runners are often categorized as forefoot, midfoot or rearfoot strikers, but how much and
why do individuals vary in foot strike patterns when running on level terrain? This study
used general linear mixed-effects models to explore both intra- and inter-individual varia-
tions in foot strike pattern among 48 Kalenjin-speaking participants from Kenya who varied
in age, sex, body mass, height, running history, and habitual use of footwear. High speed
video was used to measure lower extremity kinematics at ground contact in the sagittal
plane while participants ran down 13 meter-long tracks with three variables independently
controlled: speed, track stiffness, and step frequency. 72% of the habitually barefoot and
32% of the habitually shod participants used multiple strike types, with significantly higher
levels of foot strike variation among individuals who ran less frequently and who used lower
step frequencies. There was no effect of sex, age, height or weight on foot strike angle, but
individuals were more likely to midfoot or forefoot strike when they ran on a stiff surface, had
a high preferred stride frequency, were habitually barefoot, and had more experience run-
ning. It is hypothesized that strike type variation during running, including a more frequent
use of forefoot and midfoot strikes, used to be greater before the introduction of cushioned
shoes and paved surfaces.
Introduction
Runners are commonly categorized according to strike type (also known as footfall pattern),
and it is widely observed that more than 85% of habitually shod runners typically rearfoot
strike (RFS), in which the heel is the first part of the foot to contact the ground [1,2]. In con-
trast, some runners (many of them elite athletes) have been observed to forefoot strike (FFS),
in which the ball of the foot lands before the heel, or to midfoot strike (MFS), in which the heel
PLOS ONE | DOI:10.1371/journal.pone.0131354 July 8, 2015 1 / 17
OPENACCESS
Citation:Lieberman DE, Castillo ER, Otarola-Castillo
E, Sang MK, Sigei TK, Ojiambo R, et al. (2015)
Variation in Foot Strike Patterns among Habitually
Barefoot and Shod Runners in Kenya. PLoS ONE 10
(7): e0131354. doi:10.1371/journal.pone.0131354
Editor:Ramesh Balasubramaniam, University of
California, Merced, UNITED STATES
Received:January 22, 2015
Accepted:June 1, 2015
Published: July 8, 2015
Copyright: 2015 Lieberman et al. This is an open
access article distributed under the terms of the
Creative Commons Attribution License, which permits
unrestricted use, distribution, and reproduction in any
medium, provided the original author and source are
credited.
Data Availability Statement:All relevant data are
within the paper and its Supporting Information files.
The raw data used to make all the analyses with the
participants IDs removed has been uploaded on
figshare.com at: http://dx.doi.org/10.6084/m9.figshare.1375654. Due to ethical considerations high
speed videos are available upon request by
contacting Daniel E. Lieberman, Department of
Human Evolutionary Biology, Harvard University,
Cambridge MA 02138; email: [email protected].
edu.
Funding:This work was supported by the American
School of Prehistoric Research, Harvard University
http://creativecommons.org/licenses/by/4.0/http://dx.doi.org/10.6084/m9.figshare.1375654http://dx.doi.org/10.6084/m9.figshare.1375654http://dx.doi.org/10.6084/m9.figshare.1375654http://dx.doi.org/10.6084/m9.figshare.1375654http://creativecommons.org/licenses/by/4.0/http://crossmark.crossref.org/dialog/?doi=10.1371/journal.pone.0131354&domain=pdf -
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and ball of the foot land almost simultaneously [3]. In addition, numerous studies have found
that barefoot and minimally shod runners are more likely than habitually shod runners to MFS
or FFS [413]. However, some habitually barefoot individuals have been observed to primarily
RFS when they run [14], and people in minimal shoes are more likely to run with a RFS than
those who are barefoot [15].
Differences in strike patterns have led to numerous hypotheses about their relative costs
and benefits. Although FFS and RFS landings do not differ in terms of economy [1619], FFS
and some MFS landings differ from RFS landings in generating no discernible impact peak in
the vertical ground reaction force just after contact. Whether the rate of loading and magnitude
of impact peaks contribute to repetitive stress injuries is debated [ 2023], but impact peaks can
be uncomfortable, often causing barefoot runners to avoid RFS landings on hard surfaces with-
out a cushioned shoe [710,2023].
Regardless of the advantages and disadvantages of FFS, MFS and RFS landings, one issue
that has been insufficiently considered is variation, both within and between individuals. How
much do runners vary their strike patterns, and what causes this variation? Although runners
tend to be characterized as either rearfoot, midfoot or forefoot strikers, it is likely that most use
all three kinds of strikes but in different proportions and contexts. All people FFS when run-
ning up a steep incline, and the tendency to RFS is often greater when descending [ 24,25]. Inaddition, runners are more likely to MFS or FFS as they increase speed [ 26]. Additional factors
that may affect strike type include training and skill, fatigue, the presence of shoes, shoe design,
and substrate characteristics such as stiffness, slipperiness, unevenness and roughness. For
example, habitually shod people who normally RFS typically switch to a FFS when asked to run
barefoot on hard surfaces such as asphalt, but often continue to RFS when running barefoot on
less stiff surfaces such as grass or cushioned mats [79]. Evidence that minimally shod individ-
uals are more than twice as likely to RFS as barefoot individuals [ 79,15] suggests that sensory
feedback from the foot strongly influences strike type.
The goal of this study therefore was to explore how much runners vary strike type on level
surfaces, and to test some of the factors that may contribute to this variation. Conceptually, the
factors that influence strike type variation can be classified into three non-mutually exclusive
categories:intrinsic,extrinsicandacquired. Intrinsic factors relevant to strike type are charac-teristics of the runner that are not under control such as height, sex, age, and body mass. The
dominant extrinsic factors relevant to strike type are characteristics of a runner s environment
that potentially affect kinematics such as the nature of the substrate (e.g., surface stiffness,
slope, unevenness, slipperiness) as well as footwear characteristics such as heel cushioning that
may affect how the body interacts with the ground. Speed can also be an extrinsic factor
depending on circumstance (e.g., when a runner is required to run faster or slower, as in this
experiment). Finally, acquired factors are characteristics that a runner develops or learns. Some
acquired characteristics, such as running history, footwear history, physical fitness, strength,
and existing injuries, are often a product of an individuals background. Others such as step fre
quency may be modifiable characteristicsskillsthat are acquired through cultural processes
such as coaching, imitation, practice, and experimentation [27].
Using this conceptual framework, we tested two general hypotheses about intra- and inter-
individual strike type variation among a diverse sample of individuals who varied in several
intrinsic, extrinsic and acquired characteristics, and for whom we experimentally modified sev-
eral extrinsic and acquired variables including surface stiffness, speed, and step frequency. The
first general hypothesis (H1) is that extrinsic, intrinsic, and acquired factors influence the
degree of intra-individual variation in strike type. Specifically, because shoes slow the rate of
impact loading, limit exteroreception, and mitigate the effects of substrate variations on the
foot and the rest of the body, H1a predicts that individuals who are barefoot will use more
Foot Strike Variationamong Habitually Barefoot andShod Runners
PLOS ONE | DOI:10.1371/journal.pone.0131354 July 8, 2015 2 / 17
(DEL). The funders had no role in study design, data
collection and analysis, decision to publish, or
preparation of the manuscript.
Competing Interests:The authors have declared
that no competing interests exist.
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varied foot strike patterns than individuals who are shod. In addition, because speed and surface
stiffness may affect aspects of kinematics and kinetics relevant to strike [79,26], H1b predicts
that runners are likely to use more varied strike patterns on soft surfaces and at slower speeds.
The second general hypothesis (H2) is that a combination of extrinsic, intrinsic, and
acquired factors are predictive of foot strike angle as well as strike type both within and
between individuals. Specifically, we predict that runners are more likely to FFS as they increase
speed, increase step frequency, and run on harder surfaces. In addition, because impact peaks
can cause discomfort and might contribute to overuse injuries, especially in unshod individu-
als, we predict that runners who are habitually barefoot and run more regularly over longer dis-
tances are more likely to FFS independent of other intrinsic factors such as sex, age, and body
shape and size variation.
Materials and Methods
Study Design
Although kinematic variables such as foot strike are often compared between groups that differ
in terms of footwear use (e.g., [48,11,12,15]), the hypotheses this study tests require a com-
bined within- and between- subjects experimental design. In particular, we asked subjects whovaried in terms of the intrinsic and acquired factors described above to perform a set of trials
that independently varied three external factors: speed, surface stiffness and step frequency.
Although this study design requires repeated measurements, which can be accounted for statis-
tically using General Linear Mixed Models [36], it avoids potential sampling problems, such as
heterogeneity within and between groups as well as assignment bias.
Participants
Because this study explores both intra- and inter-individual variation, it is necessary to test the
above hypotheses with an appropriate population that varies considerably in a range of extrin-
sic, intrinsic, and acquired factors including footwear use. Almost all people in developed
nations are habitually shod, and although barefoot running has recently gained popularity incountries such as the US, few if any of these barefoot enthusiasts grew up unshod, and some
may have consciously adopted a running form advocated by books or websites. At the other
end of the continuum, most habitually barefoot populations do not practice much long dis-
tance running. For this reason, we chose to focus on Kalenjin-speaking communities from
Western part of Kenya, an area of special relevance for the questions posed by this study
because of the considerable variation in footwear usage and running habits within this popula-
tion, which includes many of the worlds best distance runners, most of whom grew up bare-
foot [27,28].
48 Kalenjin individuals (Table 1) were recruited from the region around Eldoret in the
Uasin Gishu and Nandi Counties of Kenya. 38 participants (19 M, 19 F) were adolescents
Table 1. Levene's Test of unequal variance for nominal comparisons of foot strike angle (FSA).
Comparison F-ratio p-value
Sex (male vs female) 0.1349 0.7151
Footwear (bare vs shod) 2.5124 0.1197
Surface (hard vs soft) 6.1117 0.0152
Habitually barefoot 0.1062 0.7458
Habitually shod 0.081 0.7775
doi:10.1371/journal.pone.0131354.t001
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between the ages of 13 and 17 from three schools. 19 students (10 male, 9 female) aged 1317
attended a school in a rural part of the Nandi South District where almost all the students are
primarily barefoot and very physically active [28]. The school is not directly accessible by road,
and these students walk or run barefoot an average of 7.5 3.0 km/day to travel to and from
school [29]. A few of these students wear shoes a few hours a week when they attend church
and other special occasions, but they are otherwise almost always barefoot (see below). We also
recruited 9 female students aged 1317 from a girls secondary school in Kobujoi, Kenya, and
10 male students aged 1416 from a boys secondary school in Eldoret, Kenya. These students
board at school and wear thick-soled leather shoes for most of the day, and either rubber sports
shoes (plimsoles) or cushioned athletic shoes (trainers) during athletic activities. Finally, we
recruited 10 habitually barefoot, male adults aged 2360 from the Nandi South District, Kenya.
These men walk long distances regularly, some still run several kilometers per week, and most
of them ran long distances when they were younger.
Individuals who had current lower extremity injuries or evident illness were excluded. In
order to avoid biased samples in terms of fitness, we asked the teachers at the three schools to
select only students who were averagein terms of sports ability, thus excluding participants
who either exceptional or poor in athletics.
Ethics Statement
Approval for the human experimental study described in this paper was granted by the Har-
vard University Committee on the Use of Human Subjects (protocol F23121), and by the Moi
University Medical Institutional Research And Ethics Committee (protocol 00695). As
approved by the aforementioned committees, written informed consent for minors was pro-
vided by their teachers; informed consent was provided orally by adults who were unable to
read and documented with their signature.
Anthropometrics and Background Information
Basic anthropometrics were collected from all participants including height, body mass, and
leg length (from the greater trochanter to the base of the heel). An orthopedic doctor (POM)examined all participants for lower extremity injuries. All participants (some of whom were
not literate) were asked how far they walk and run on average each day, their regular physical
activities, and what kinds of footwear they use. All questions were asked on two different occa-
sions, either in Kalenjin or Kiswahili; one of the questioners (MS) speaks Kalenjin, knows the
region intimately, and was able to evaluate how far each participant had to walk or run every
day. Answers were then averaged. Since footwear usage and running history could not be quan
tified precisely as continuous variables, answers to these questions were binned into four rank
order categories. Footwear score categories were: 1, almost always shod (less than 10% outdoor
activity spent barefoot or in minimal shoes); 2, usually shod (mostly wear shoes, but do sports
either barefoot or in minimal shoes); 3, mixed (sometimes walk, run or do physical activity in
normal shoes and sometimes barefoot or in minimal shoes); 4, mostly barefoot (more than
80% of walking, running and physical activity done either barefoot or in minimal shoes). Run-ning history categories were: 1, little (run less than 5 km/week); 2, occasional (run 5 10 km a
week on an occasional but non-regular basis; 3, moderate (run 510 km a week on a regular
basis); high (run >10 km a week on a regular basis).
Experimental Trials
Participants were asked to wear whatever footwear they normally use (if applicable), and to
wear shorts or skirts that could be rolled above the knee. In order to record 2-dimensional
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kinematics in lateral view, reflective tape markers were placed on the following locations on
one side of the body: the greater trochanter, the center of the knee (in between the lateral femo-
ral epicondyle and the lateral tibial plateau), the lateral malleolus, the lateral surface of the 5 th
metatarsal head, and the lateral tuber calcaneus. Participants were then photographed with a
visual scale in lateral and frontal position with a numeric identification. All participants were
then instructed to run around an open field at a pace they would choose if running a long dis-
tancefor approximately 5 minutes, at which point step frequency was then measured using an
adjustable metronome (Matrix, New Market, VA, USA) fitted with an earpiece. Preferred step
frequency (PSF) was recorded only for step frequencies that did not deviate by more than 4
steps/minute over a minimum of 30 seconds.
After warm-up, each participants kinematics was immediately recorded in lateral view on
two adjacent tracks approximately 1315 m in length. The hardtrack was the unaltered,
grass-free, compact surface of a field, similar to the stiffness of a dirt road s surface, and typical
of the surfaces on which the participants normally run when traveling or doing athletics. A
softtrack was excavated parallel to the hard track by digging down 10 cm with a pickaxe,
tamping down the earth, and then raking the dirt to create a smooth, soft surface. Penetrometer
measurements repeated on each track (AMS Corp, American Falls, ID) indicate that the aver-
age compression strength of the hard track (3.85 kg/cm2
0.29 S.D.) was 5.5 times greater thanthe soft track (0.70 kg/cm20.27 S.D.). The soft track was raked between each set of trials, and
re-excavated regularly to maintain a similar compliant surface for all participants. Small flags
were used to mark the borders of the two tracks. A high-speed video camera (Casio EX-ZR100)
was positioned at 0.7 m height approximately 4 m lateral to the 10 m point on the track, pro-
viding an additional 35 m of track beyond the field of the camera. All sequences were
recorded at 240 frames per second.
For each trial, participants were asked to run down the track while looking forward and
without decelerating until they had passed a marker positioned approximately 3 meters beyond
the cameras field of view. Participants were asked to run down both the hard and soft tracks at
approximately 3.0 m/sec (slow) and 4.0 m/sec (fast) at several step frequencies: the previ-
ously determined preferred step frequency (PSF), and at 150, 170 and 190 steps/min. As a
result, each participant ran a minimum of 16 conditions. Step frequency was controlled using asmall, lightweight digital metronome either handheld or clipped onto clothing (Seiko DM50,
SeikoUSA, Mahwah, NJ). For each trial, the participant was familiarized with the frequency
and then asked to try to maintain that frequency for the entire length of the track. There was
no landing target on the track in order to avoid having participants alter their gaits by either
shortening or lengthening their steps as they passed the camera s field of view. If the marked
foot did not land in front of the camera, the trial was repeated without explaining the reason
for repetition until a minimum of two trials were recorded for each speed, step frequency, and
track. Following these trials, we administered the mile run test to the adolescent participants
from each school according to methods outlined by the FITNESSGRAM test [30]. To avoid
influencing how participants ran, we asked no questions about running form before or after
the trials, and neither the participants nor their teachers were informed of the experiment s
objectives.
Kinematic Analysis
All video sequences were converted to stacks of TIFF files and analyzed using ImageJ, version
1.46r (http://imagej.nih.gov/ij). A visual scale was determined for each participant using the
measured distance between the lateral malleolus and knee markers. Since running speed was
not controlled precisely during the experiment, running speed for each trial was quantified by
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measuring the horizontal translation of the marker on the greater trochanter between two
homologous points during a stride cycle (e.g., toe-off to toe-off, or foot strike to foot strike for
the same foot) relative to time (calculated from the number of frames divided by frame rate).
Foot strike was measured using only high-speed sequences in which the marked foot landed
in front of the camera permitting a clear view of the foot s lateral margin, which has been
shown to yield high accuracy and reliability [31]. Foot strike angle (FSA) was quantified as a
continuous variable by measuring the orientation of the calcaneus and 5 th metatarsal head
markers relative to horizontal at the first frame of contact minus the same angle measured at
foot flat [7]. Since FSA is a continuous variable but foot strike itself is a nominal variable, strike
types were also classified using the following criteria: FFS, angles above 0.3; MFS, angles
between 0.3 and -5.6; RFS, angles below -5.6. In order to avoid classifying RFS and FFS land-
ings as MFS landings, these cutoff values are more conservative than those used by Altman and
Davis [31]. The correlation between strike type and FSA was 0.95 (p
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participants. To account for repeated measures, these tests used the mean variance of each indi-
vidual. A Chi-squared analysis was also used to test if the proportion of individuals who varied
their strike type differed between the barefoot and shod participants. Second, to test if there is a
relationship between levels of variation in foot strike and intrinsic, extrinsic and acquired fac-
tors that are continuously distributed, we used a bivariate General Linear Mixed Model
(GLMM) to calculate the residuals of the regression between FSA and each predictor variable
using a subject identifier as the random effect to account for the non-independent error gener-
ated by repeated measures on the same individuals [32]. We then used a second GLMM to
regress the absolute value of these residuals against the relevant predictor variable. A slope (the
coefficient of the GLMM) significantly different from zero indicates a significant increase or
decrease in variation with respect to the predictor variable. Since GLMMs assume that variables
are normally distributed and in comparable units, non-normally distributed variables (assessed
using a Shapiro-Wilk test) were log-transformed, and then all variables were converted to Z-
scores.
In order to test Hypothesis 2, we used multivariate GLMMs to model the effects of the
intrinsic, extrinsic, and acquired variables on foot strike across treatments. In the first GLMM,
the dependent variable was FSA was regressed against the fixed effects included several vari-
ables classified as extrinsic (substrate stiffness), intrinsic (age, sex, height, body mass), andacquired factors (footwear history and running history; preferred step frequency; and the speed
at which the participants could run a mile, a proxy for overall physical fitness). The first
GLMM took the following form:
FSA b1Surface b
2Age b
3Sex b
4Height b
5Body Mass b
6Footwear History
b7Running History b
8Preferred Step Frequency b
9Mile Time ZU
A second GLMM (S1 Table) was also calculated to test the effects of kinematics on foot strike.
In this GLMM, the dependent variable was FSA and the xed effects were aspects of kinematics
(speed, step frequency, trunk angle, hip angle, knee angle, and overstride relative to the knee).
This took the following form:
FSA b1Speed b
2Step Frequency b
3Trunk Angle b
4Hip Angle b
5Knee Angle
b6Ankle Angle b
7Overstride ZU
In both models,iis thexed-effect coefcient for theith predictor,Zis the design matrix for
the random grouping variable,Uis a vector of random effects, and is residual model error. A
subject identier was used as the random grouping effect to account for repeated measures on
the same individuals. In addition, all variables were transformed to Z-scores, and non-normally
distributed variables (assessed using a Shapiro-Wilk test) were log-transformed.
A few of the variables (including the primary outcome measurement, FSA) were not nor-
mally distributed after transformation to Z-scores, and some variables are highly collinear (e.g.,
height and body mass, speed and step frequency). Therefore, to account for non-normality and
isolate the potential effects of multicollinearity on significance testing, we also used a non-
parametric residual randomization method to calculate p-values in the GLMMs. Residual ran-
domization is a type of permutation test in which statistical significance is tested by permuting
the residuals of a model rather than the observations [33]. In this study, residual randomization
is used to evaluate the significance of each variable s effect independently, removing partial
effects of other collinear variables (seeS1 File).
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Results
Because this study aimed to sample a wide range of variation both within and between different
groups, we begin with a summary of the variation sampled. In terms of external factors, average
height was 160.9 9.5 cm (range 142177) in the barefoot population and 162.38.1 cm (range
147174) in the shod population (p = 0.517); average body mass was 47.08.4 kg (range 32
62) in the barefoot population and 54.08.1 kg (range 3765) in the shod population(p = 0.038); average age was 24.815.1 years (range 1337) in the barefoot population, and
15.4 1.07 years (range 1318) in the habitually shod population. In terms of acquired variables
measured, the average footwear score in the barefoot population was 3.341.2 (range 24),
higher (p
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Foot Strike Variationamong Habitually Barefoot andShod Runners
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footwear history, running history, preferred step frequency, and the speed at which the partici-
pants could run a mile. As the coefficients (which represent the slope of the relationshipbetween FSA and each predictor variable) inTable 3indicate, none of the intrinsic variables
(sex, age, body mass) have an effect on FSA at conventional levels of significance (p
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acquired variables and FSA. AsFig 3Ashows, individuals who spend more time barefoot show
considerable variation in FSA but tend to have more positive FSAs, whereas individuals who
are more habitually shod have more negative FSAs (ANOVA, p
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individuals. This variation is highlighted by the plot of every FSA recorded in the study ( Fig 1),
which shows that the average intra-individual standard deviation of FSA was 4.12, and that
while a majority of participants (56%) used a combination of FFS, MFS and RFS landings, 72%
of the barefoot runners and 32% of the shod runners used multiple strike types.This study tested two general hypotheses regarding the effects of intrinsic, extrinsic, and
acquired factors on variations in observed in foot strike. The first general hypothesisthat cer-
tain factors influence the degree of variation in strike typewas supported (seeTable 1and
Table 2). Although none of the intrinsic factors measured (height, sex, age, and body mass)
affected the degree of variation in FSA, several extrinsic and acquired factors did influence FSA
variation. In particular, there was a significantly greater degree of FSA variability within indi-
viduals who used lower step frequencies and who typically ran less. In addition, although FSA
Fig 2.Sources of variation in foot strikeangle (FSA). a) Regression of speed versusFSA; b) regression of measured step frequencyversus FSA; c)regression of preferred stride frequencyversus FSA; d) Box (standarderror)and whisker (standard deviation) plot of difference in FSA on hard versussofttracksfor habitually barefoot and shod individuals (more positive valuesindicatemore dorsiflexed FSA on soft surface; more negative valuesindicate moreplantar flexedFSA on soft surface); x marks indicate maximum andminimum values.
doi:10.1371/journal.pone.0131354.g002
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variation was not affected by footwear history, individuals who were barefoot had significantly
more variable foot strike types than those who were wearing shoes. The explanation for this
seemingly contradictory result is that average FSA among barefoot individuals was 1.11 5.3,
whereas the average FSA among those who were habitually shod was -8.3 6.1 (t-test,
p
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would also be more likely to MFS or FSS. These hypotheses were all supported. In particular,
the GLMM (Table 3) revealed significant effects of track stiffness, preferred step frequency,
footwear history, running history and mile time speeds. Put simply, individuals were less likely
to RFS when they ran on a harder track (Fig 2D), preferred higher step frequencies (Fig 2C),
were able to run faster, were experienced runners (Fig 3A), and were habitually barefoot (Fig
3B). In contrast, there was no effect on FSA from age, sex, body mass, height, or speed ( Fig
2A).
These results are consistent with a previous, smaller comparison of barefoot and shod
Kalenjin individuals that sampled a more limited range of faster speeds [7], as well as studies
that compare running form among populations that vary in footwear use [ 11,15] or in which
habitually shod individuals have been studied both barefoot and shod [8,9,34]. Although bare-
foot individuals sometimes RFS, they are more likely to FFS and MFS depending on conditions
and experience; in contrast, habitually shod individuals are more likely to RFS under a range of
conditions.
Although not a focus of this study, the results presented here confirm those of previous
studies that compared kinematics and kinetics between barefoot and shod runners [ 412,15].
In general, the habitually barefoot participants landed with more flexed knees and hips, they
had slightly more vertical trunks, they preferred higher step frequencies, and they were lesslikely to overstride (S2 Table). When a GLMM was used to tease apart which of these variables
were associated with variations in FSA, the strongest predictor was ankle angle, with significant
associations also evident for overstride and trunk angle (S1 Table).
Before considering the implications of these results, it is worth summarizing the studys lim
itations. One problem is the limited range of subjects, conditions, and factors sampled. We
were unable to include adult women, and the sample sizes for each group were necessarily lim-
ited by time and opportunity. Broadening the sample in terms of age, sex, running experience,
and footwear history would likely reveal additional variability. In addition, the experimental
design did not look at fatigue, which can increase the likelihood of using a RFS [ 35], and only a
few external factors hypothesized to influence kinematics (notably speed, step frequency and
surface stiffness) were manipulated. Future studies would benefit from examining substrate
factors such as slipperiness, smoothness, inclines, and changes in direction of the sort that run-ners encounter when they run on trails and other variable environments that, until relatively
recently, were the primary contexts in which people ran. Another necessary limitation of the
study was to measure only sagittal plane kinematics using video without collecting information
on ground reaction forces and muscle function. A final concern was the participants ability to
run normally. Although the experiment was not conducted in the laboratory on a treadmill,
running at different speeds on a track with markers taped to one s joints while trying to adapt
ones step frequency to a metronome is an unusual experience that can interfere with normal
running form. This concern, however, applies to all studies of running kinematics and kinetics,
and it is arguable that the conditions tested here are a step in the direction of understanding
variability in running form beyond the laboratory and among individuals who are not just
habitually shod from developed countries. Although such people are the focus of most
research, they are unusual from an evolutionary perspective [36].
These limitations aside, the studys results have some relevance for current discussions
about running form. Most importantly, very few studies on running biomechanics have sam-
pled runners who are not habitually shod and in the natural settings in which people used to
locomote rather than in controlled laboratory conditions, primarily on treadmills or over force
plates [7,11,13,14]. It is reasonable to hypothesize that these modern contexts limit variation in
foot strike as well as other aspects of kinematics. The results presented here raise the possibility
that for much of human evolution foot strike patterns were more variable. The two most
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obvious factors that have potentially contributed to less variation in how people run is increased
use of flat, paved surfaces and treadmills for running, and the prevalence of running shoes with
elevated, cushioned heels that have been available only since the 1970s [ 7]. Just as cushioned
heels facilitate RFS landings on hard surfaces, it is reasonable to hypothesize that the barefoot
individuals measured in this study were more likely to run with a RFS on the soft trackway
because softer substrates, like cushioned heels, make RFS landings more comfortable by lowering
the rate of loading of the impact peak [7]. Although soft and smooth surfaces no doubt existed in
the past such as along lakeshores and in sandy environments, most people typically walked and
ran on compacted soil with rocks, vegetation, and features that increase substrate complexity and
stiffness. Walking and running without shoes on these surfaces unquestionably elicits much
more varied and extreme stimuli from sensory nerves on the glabrous surface of the sole. It is
therefore reasonable to hypothesize that people ran with more varied kinematics prior to the
invention of shoes, which probably occurred in the last 40,000 years [37].
Another factor that may have affected variation in running form is skill. Since the running
boom that began in the 1970s, there has been an increase in running among amateurs, who
usually get less coaching and train less intensively than athletes who are professional or on
teams [38]. One hypothesis that merits further testing is that untrained, amateur runners in
developed nations are more likely to run like the habitually shod Kalenjin studied here, with arelatively slower step frequency and a greater proclivity to land with a dorsiflexed foot, hence a
RFS. This observation leads to the hypothesis that a contributing factor to Kalenjin excellence
in distance running might be that most elite Kalenjin runners grew up running long distances
without shoes on a regular basis in the same conditions as the habitual barefoot participants
analyzed in this study [27]. Although habitually barefoot people from the Daasanach tribe in
northern Kenya were observed to mostly run with a RFS at slow speeds (2.13.0 m/s) a possible
explanation for this different result, apart from speed, is that these individuals live in a hot,
sandy desert and do not run often much [14]. Other studies of adults from habitually barefoot
and minimally shod populations found that individuals (especially men) were more likely to
FFS or MFS [7,11,13].
Finally, what do these results mean for habitually shod individuals who run mostly on pave-
ment and treadmills, and wonder how to make sense of diverse arguments about minimalshoes, cushioning, and strike type? First, the restricted variation in strike type among habitually
shod runners today may be a recent phenomenon, and it would be useful to test if runners
adopt more variation when running on trails rather than on pavement or treadmills. In addi-
tion, although rearfoot and forefoot striking are both normal, everything involves trade-offs.
RFS landings have the potential advantages of being comfortable in shoes or on soft surfaces,
they require less calf and foot muscle strength, and they lessen external moments acting around
the ankle [39]. Their potential disadvantages are that they cause impact peaks whose rate and
magnitude are hypothesized by some researchers to be related to some repetitive stress injuries,
they increase external moments around the knee, and certain kinematic patterns associated
with (but not exclusive to) RFS gaits, such as overstriding and extended knees at landing, are
implicated in some repetitive stress injuries [40]. More research is needed to evaluate the costs
and benefits of different strike types, but one hypothesis that also needs to be explored is that
running with more kinematic variation, as perhaps occurs during trail running, is more natural
and may also be beneficial.
Supporting Information
S1 File. Residual Randomization Methods.
(DOCX)
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S1 Table. GLMM analysis of effects of kinematic variables on Foot strike Angle (FSA).
(DOCX)
S2 Table. Group means and standard deviations for major variables studied.
(DOCX)
Acknowledgments
This research was funded by Harvard University. We thank H. Dingwall, E. Hintze, E. Maritim
N. Keter, E. Anjila, and the teachers and students at the Pemja, AIC Chebisaas, and Mothers of
Apostles Schools, Kenya.
Author Contributions
Conceived and designed the experiments: DEL ERC. Performed the experiments: DEL ERC
MKS TKS PO. Analyzed the data: DEL ERC EOC. Wrote the paper: DEL ERC EOC TKS MKS
RO PO YP.
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